Effects of seawater acidification on a coral reef meiofauna community

Despite the increasing risk that ocean acidification will modify benthic communities, great uncertainty remains about how this impact will affect the lower trophic levels, such as members of the meiofauna. A mesocosm experiment was conducted to investigate the effects of water acidification on a phytal meiofauna community from a coral reef. Community samples collected from the coral reef subtidal zone (Recife de Fora Municipal Marine Park, Porto Seguro, Bahia, Brazil), using artificial substrate units, were exposed to a control pH (ambient seawater) and to three levels of seawater acidification (pH reductions of 0.3, 0.6, and 0.9 units below ambient) and collected after 15 and 30 d. After 30 d of exposure, major changes in the structure of the meiofauna community were observed in response to reduced pH. The major meiofauna groups showed divergent responses to acidification. Harpacticoida and Polychaeta densities did not show significant differences due to pH. Nematoda, Ostracoda, Turbellaria, and Tardigrada exhibited their highest densities in low-pH treatments (especially at the pH reduction of 0.6 units, pH 7.5), while harpacticoid nauplii were strongly negatively affected by low pH. This community-based mesocosm study supports previous suggestions that ocean acidification induces important changes in the structure of marine benthic communities. Considering the importance of meiofauna in the food web of coral reef ecosystems, the results presented here demonstrate that the trophic functioning of coral reefs is seriously threatened by ocean acidification.     

An introduction to the biogeochemical cycling of calcium and substitutive strontium in living coral reef mesocosms

The coral reef mesocosms designed by the Smithsonian Institution's Dr. Walter Adey, his Marine System Laboratory personnel, and staff members of the Pittsburgh Aqua-Zoo simulate most of the physical, chemical, and biological parameters found in natural Caribbean coral reefs. After developing the mesocosm in Pittsburgh, an evaluation and comparison between natural reef seawater sources and closed mesocosm seawater conditions indicated that an additional parameter should be investigated. It was hypothesized that, given time, the aragonite- and calcite (CaCO₃ crystal forms)-producing organisms in the closed mesocosms could deplete the seawater of available Ca²⁺ and substitutive Sr²⁺. Atomic absorption spectrophotometry was utilized to determine concentrations in the seawater over time. Results showed a substantial reduction in dissolved Ca and Sr in the mesocosm after approximately two years. Dissolved aragonitic Halimeda algae parts were put into the system for replacive purposes. In terms of the biogeochemical cycling of Ca²⁺ and Sr²⁺, the coral reef mesocosm organisms behaved similarly to natural reefs, which have a constant supply of dissolved Ca²⁺ and Sr²⁺. Further research utilizing radiolabeled sources of Ca²⁺, Sr²⁺, and Mg²⁺, in conjunction with in vivo scanning electron microscopy (SEM) and growth increment studies, are recommended for determining the exact biogeochemical pathways for these elements in coral reefs, and to quantify growth parameters. © 1993 Wiley-Liss, Inc.     

Near-future ocean acidification enhances the feeding rate and development of the herbivorous juveniles of the crown-of-thorns starfish, <Emphasis Type="Italic">Acanthaster planci</Emphasis>

Population outbreaks of the corallivorous crown-of-thorns starfish, Acanthaster planci, are a major contributor to the decline in coral reef across the Indo-Pacific. The success of A. planci and other reef species in a changing ocean will be influenced by juvenile performance because the naturally high mortality experienced at this sensitive life history stage maybe exacerbated by ocean warming and acidification. We investigated the effects of increased temperature and acidification on growth of newly metamorphosed juvenile A. planci and their feeding rates on crustose coralline algae (CCA) during the initial herbivorous phase of their life history. The juveniles were exposed to three temperature (26, 28, 30 °C) and three pH (NIST scale: 8.1, 7.8, 7.6) levels in a flow-through cross-factorial experiment. There were positive but independent effects of warming and acidification on juvenile growth and feeding. Early juveniles were highly tolerant to moderate increases in temperature (+2 °C above ambient) with the highest growth at 30 °C. Growth and feeding rates of A. planci on CCA were highest at pH 7.6. Thus, ocean warming and acidification may enhance the success of A. planci juveniles. In contrast to its coral prey, at this vulnerable developmental stage, A. planci appears to be highly resilient to future ocean change. Success of juveniles in a future ocean may have carry-over effects into the coral-eating life stage, increasing the threat to coral reef systems.     

Combined effects of warming and ocean acidification on coral reef Foraminifera Marginopora vertebralis and Heterostegina depressa

Warming and changes in ocean carbonate chemistry alter marine coastal ecosystems at an accelerating pace. The interaction between these stressors has been the subject of recent studies on reef organisms such as corals, bryozoa, molluscs, and crustose coralline algae. Here we investigated the combined effects of elevated sea surface temperatures and pCO₂ on two species of photosymbiont-bearing coral reef Foraminifera: Heterostegina depressa (hosting diatoms) and Marginopora vertebralis (hosting dinoflagellates). The effects of single and combined stressors were studied by monitoring survivorship, growth, and physiological parameters, such as respiration, photochemistry (pulse amplitude modulation fluorometry and oxygen production), and chl a content. Specimens were exposed in flow-through aquaria for up to seven weeks to combinations of two pCO₂ (~790 and ~490 µatm) and two temperature (28 and 31 °C) regimes. Elevated temperature had negative effects on the physiology of both species. Elevated pCO₂ had negative effects on growth and apparent photosynthetic rate in H.depressa but a positive effect on effective quantum yield. With increasing pCO₂, chl a content decreased in H. depressa and increased in M. vertebralis. The strongest stress responses were observed when the two stressors acted in combination. An interaction term was statistically significant in half of the measured parameters. Further exploration revealed that 75 % of these cases showed a synergistic (= larger than additive) interaction between the two stressors. These results indicate that negative physiological effects on photosymbiont-bearing coral reef Foraminifera are likely to be stronger under simultaneous acidification and temperature rise than what would be expected from the effect of each of the stressors individually.     

Coral reefs modify their seawater carbon chemistry – implications for impacts of ocean acidification

Reviews suggest that that the biogeochemical threshold for sustained coral reef growth will be reached during this century due to ocean acidification caused by increased uptake of atmospheric CO₂. Projections of ocean acidification, however, are based on air‐sea fluxes in the open ocean, and not for shallow‐water systems such as coral reefs. Like the open ocean, reef waters are subject to the chemical forcing of increasing atmospheric pCO₂. However, for reefs with long water residence times, we illustrate that benthic carbon fluxes can drive spatial variation in pH, pCO₂ and aragonite saturation state (Ω_a) that can mask the effects of ocean acidification in some downstream habitats. We use a carbon flux model for photosynthesis, respiration, calcification and dissolution coupled with Lagrangian transport to examine how key groups of calcifiers (zooxanthellate corals) and primary producers (macroalgae) on coral reefs contribute to changes in the seawater carbonate system as a function of water residence time. Analyses based on flume data showed that the carbon fluxes of corals and macroalgae drive Ω_ain opposing directions. Areas dominated by corals elevate pCO₂ and reduce Ω_a, thereby compounding ocean acidification effects in downstream habitats, whereas algal beds draw CO₂ down and elevate Ω_a, potentially offsetting ocean acidification impacts at the local scale. Simulations for two CO₂ scenarios (600 and 900 ppm CO₂) suggested that a potential shift from coral to algal abundance under ocean acidification can lead to improved conditions for calcification in downstream habitats, depending on reef size, water residence time and circulation patterns. Although the carbon fluxes of benthic reef communities cannot significantly counter changes in carbon chemistry at the scale of oceans, they provide a significant mechanism of buffering ocean acidification impacts at the scale of habitat to reef.     

Dissolved inorganic carbon and total alkalinity of a Hawaiian fringing reef: chemical techniques for monitoring the effects of ocean acidification on coral reefs

There is an interest in developing approaches to “ecosystem-based” management for coral reefs. One aspect of ecosystem performance is to monitor carbon metabolism of whole communities. In an effort to explore robust techniques to monitor the metabolism of fringing reefs, especially considering the possible effects of ocean acidification, a yearlong study of the carbonate chemistry of a nearshore fringing reef in Hawaii was conducted. Diurnal changes in seawater carbonate chemistry were measured once a week in an algal-dominated and a coral-dominated reef flat on the Waimanalo fringing reef, Hawaii, from April of 2010 until May of 2011. Calculated rates of gross primary production (GPP) and net community calcification (G) were similar to previous estimates of community metabolism for other coral reefs (GPP 971 mmol C m^?2 d^?1; G 186 mmol CaCO₃ m^?2 d^?1) and indicated that this reef was balanced in terms of organic metabolism, exhibited net calcification, and was a net source of CO₂ to the atmosphere. Average slopes of total alkalinity versus dissolved inorganic carbon (TA–DIC slope) for the coral-dominated reef flat exhibited a greater calcification-to-net photosynthesis ratio than for the algal-dominated reef flat (coral slope vs. algal slope). Over the course of the time series, TA–DIC slopes remained significantly different between sites and were not correlated with diurnal averages in reef-water residence time or solar irradiance. These characteristic slopes for each reef flat reflect the relationship between carbon and carbonate community metabolism and can be used as a tool to monitor ecosystem function in response to ocean acidification.     

Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.     

Larvae of the coral eating crown‐of‐thorns starfish,Acanthaster planci in a warmer‐high CO2 ocean

Outbreaks of crown‐of‐thorns starfish (COTS), Acanthaster planci, contribute to major declines of coral reef ecosystems throughout the Indo‐Pacific. As the oceans warm and decrease in pH due to increased anthropogenic CO₂ production_, coral reefs are also susceptible to bleaching, disease and reduced calcification. The impacts of ocean acidification and warming may be exacerbated by COTS predation, but it is not known how this major predator will fare in a changing ocean. Because larval success is a key driver of population outbreaks, we investigated the sensitivities of larval A. planci to increased temperature (2–4 °C above ambient) and acidification (0.3–0.5 pH units below ambient) in flow‐through cross‐factorial experiments (3 temperature × 3 pH/pCO₂ levels). There was no effect of increased temperature or acidification on fertilization or very early development. Larvae reared in the optimal temperature (28 °C) were the largest across all pH treatments. Development to advanced larva was negatively affected by the high temperature treatment (30 °C) and by both experimental pH levels (pH 7.6, 7.8). Thus, planktonic life stages of A. planci may be negatively impacted by near‐future global change. Increased temperature and reduced pH had an additive negative effect on reducing larval size. The 30 °C treatment exceeded larval tolerance regardless of pH. As 30 °C sea surface temperatures may become the norm in low latitude tropical regions, poleward migration of A. planci may be expected as they follow optimal isotherms. In the absence of acclimation or adaptation, declines in low latitude populations may occur. Poleward migration will be facilitated by strong western boundary currents, with possible negative flow‐on effects on high latitude coral reefs. The contrasting responses of the larvae of A. planci and those of its coral prey to ocean acidification and warming are considered in context with potential future change in tropical reef ecosystems.     

The reef-building coral Siderastrea siderea exhibits parabolic responses to ocean acidification and warming

Anthropogenic increases in atmospheric CO₂ over this century are predicted to cause global average surface ocean pH to decline by 0.1–0.3 pH units and sea surface temperature to increase by 1–4°C. We conducted controlled laboratory experiments to investigate the impacts of CO₂-induced ocean acidification (pCO₂ = 324, 477, 604, 2553 µatm) and warming (25, 28, 32°C) on the calcification rate of the zooxanthellate scleractinian coral Siderastrea siderea, a widespread, abundant and keystone reef-builder in the Caribbean Sea. We show that both acidification and warming cause a parabolic response in the calcification rate within this coral species. Moderate increases in pCO₂ and warming, relative to near-present-day values, enhanced coral calcification, with calcification rates declining under the highest pCO₂ and thermal conditions. Equivalent responses to acidification and warming were exhibited by colonies across reef zones and the parabolic nature of the corals'' response to these stressors was evident across all three of the experiment''s 30-day observational intervals. Furthermore, the warming projected by the Intergovernmental Panel on Climate Change for the end of the twenty-first century caused a fivefold decrease in the rate of coral calcification, while the acidification projected for the same interval had no statistically significant impact on the calcification rate—suggesting that ocean warming poses a more immediate threat than acidification for this important coral species.     

First experimental evidence of corals feeding on seagrass matter

We present the first experimental evidence of a coral (Oulastrea crispata) ingesting and assimilating seagrass material. Tropical seagrass meadows export a substantial portion of their productivity and can provide an important source of nutrients to neighbouring systems such as coral reefs; however, little is known about the mechanisms of this link. To investigate whether seagrass nutrient uptake via coral heterotrophy is possible, we conducted a feeding experiment with seagrass particulate and dissolved organic matter. Using gut extractions and stable isotope analyses, we determined that O. crispata ingested ¹⁵N-enriched seagrass particles and assimilated the nitrogen into its tissue at a rate of 0.75 μg N cm^?2 h^?1. Corals took up nitrogen from dissolved matter at a comparable rate of 0.98 μg N cm^?2 h^?1. While other ecological connections between seagrass meadows and reef ecosystems are well known, our results suggest a previously unstudied direct nutritional link between seagrasses and corals.     

Structural and physiological responses of <Emphasis Type="Italic">Halodule wrightii</Emphasis> to ocean acidification

Coastal areas face high variability of seawater pH. Ocean acidification (OA) and local stressors are enhancing this variability, which poses a threat to marine life. However, these organisms present potential phenotypic plasticity that can offer physiological and structural tools to survive in these extreme conditions. In this study, we evaluated the effects of elevated CO₂ levels and consequent pH reduction on the physiology, anatomy and ultrastructure of the seagrass Halodule wrightii. A mesocosm study was conducted in an open system during a 30-day experiment, where different concentrations of CO₂ were simulated following the natural variability observed in coastal reef systems. This resulted in four experimental conditions simulating the (i) environmental pH (control condition, without CO₂ addition) and (ii) reduced pH by ? 0.3 units, (iii) ? 0.6 units and (iv) ? 0.9 units, in relation to the field condition. The evaluated population only suffered reduced optimum quantum yield (Y(II)), leaf width and cross-section area under the lowest CO₂ addition (? 0.3 pH units) after 30 days of experiment. This fitness commitment should be related to carbon concentration mechanisms present in the evaluated species. For the highest CO₂ level, H. wrightii demonstrated a capacity to compensate any negative effect of the lowest pH. Our results suggest that the physiological behaviour of this primary producer is driven by the interactions among OA and environmental factors, like irradiance and nutrient availability. The observed behaviour highlights that high-frequency pH variability and multifactorial approaches should be applied, and when investigating the impact of OA, factors like irradiance, nutrient availability and temperature must be considered as well.     

Positive genetic associations among fitness traits support evolvability of a reef‐building coral under multiple stressors

Climate change threatens organisms in a variety of interactive ways that requires simultaneous adaptation of multiple traits. Predicting evolutionary responses requires an understanding of the potential for interactions among stressors and the genetic variance and covariance among fitness‐related traits that may reinforce or constrain an adaptive response. Here we investigate the capacity of Acropora millepora, a reef‐building coral, to adapt to multiple environmental stressors: rising sea surface temperature, ocean acidification, and increased prevalence of infectious diseases. We measured growth rates (weight gain), coral color (a proxy for Symbiodiniaceae density), and survival, in addition to nine physiological indicators of coral and algal health in 40 coral genets exposed to each of these three stressors singly and combined. Individual stressors resulted in predicted responses (e.g., corals developed lesions after bacterial challenge and bleached under thermal stress). However, corals did not suffer substantially more when all three stressors were combined. Nor were trade‐offs observed between tolerances to different stressors; instead, individuals performing well under one stressor also tended to perform well under every other stressor. An analysis of genetic correlations between traits revealed positive covariances, suggesting that selection to multiple stressors will reinforce rather than constrain the simultaneous evolution of traits related to holobiont health (e.g., weight gain and algal density). These findings support the potential for rapid coral adaptation under climate change and emphasize the importance of accounting for corals’ adaptive capacity when predicting the future of coral reefs.     

Heterogeneous dissolved organic nitrogen supply over a coral reef: first evidence from nitrogen stable isotope ratios

Dissolved organic nitrogen (DON) potentially plays a major role in sustaining the high productivity and biological diversity of coral reefs. However, data are scarce regarding sources and sinks of DON. This study, for the first time, determined the ¹⁵N isotopic composition of total dissolved nitrogen (δ¹⁵N_TDN), reflecting the isotopic signature of DON, in the water column over a coral reef. The uniformity in δ¹⁵N_TDN during high tide (3.2 ± 0.3 ‰) indicated that the DON was mainly derived from offshore waters. In contrast, higher spatial heterogeneity of δ¹⁵N_TDN (3.1 ± 0.9 ‰) and DON concentrations during low tide indicated the existence of local DON sources patchily distributed over the reef. Low δ¹⁵N_TDN values located mid-reef were indicative of DON release from organisms that obtained their N via N₂ fixation, whereas high δ¹⁵N_TDN appeared to reflect localized release of DON by organisms exposed to dissolved inorganic nitrogen with elevated ¹⁵N, such as from terrestrial and offshore inputs. Collectively, the results highlight the importance of spatial patterns in DON release from reef communities in the N cycling of coral reefs.     

Community metabolism of a coral reef exposed to naturally varying dissolved inorganic nutrient loads

Daily community rates of calcification, photosynthesis and respiration were measured on a coral reef located in the Northern Red Sea, Gulf of Eilat, Israel between March 2000 and March 2002. This reef is exposed to seasonally varying levels of inorganic nutrient loading due to mixing and stratification of the adjacent open sea water column. Net production measurements were positively and linearly correlated with open sea nutrient levels, and the community photosynthesis to respiration ratio varied between 0.9 and 1.7 accordingly. Community calcification varied between 30 ± 20 and 60 ± 20 mmol C m⁻² day⁻¹ during summer and winter, respectively. Under increased nutrient loading the relation between community calcification and aragonite saturation state is suppressed by 30% on average. Both of these findings demonstrate the deleterious effects of nutrient loading on coral reefs.     

Ocean acidification accelerates net calcium carbonate loss in a coral rubble community

Coral rubble communities are an important yet often overlooked component of a healthy reef ecosystem. The organisms inhabiting reef rubble are primarily bioeroders that contribute to the breakdown and dissolution of carbonate material. While the effects of ocean acidification on calcifying communities have been well studied, there are few studies investigating the response of bioeroding communities to future changes in pH and calcium carbonate saturation state. Using a flow-through pH-stat system, coral rubble pieces with a naturally occurring suite of organisms, along with bleached control rubble pieces, were subjected to three different levels of acidification over an 8-week period. Rates of net carbonate loss in bleached control rubble doubled in the acidification treatments (0.02 vs. 0.04% CaCO₃ d^?1 in ambient vs. moderate and high acidification), and living rubble communities experienced significantly increased rates of net carbonate loss from ambient to high acidification conditions (0.06 vs. 0.10% CaCO₃ d^?1, respectively). Although more experimentation is necessary to understand the long-term response and succession of coral rubble communities under projected conditions, these results suggest that rates of carbonate loss will increase in coral rubble as pH and calcium carbonate saturation states are reduced. This study demonstrates a need to thoroughly investigate the contribution of coral rubble to the overall carbonate budget, reef resilience, recovery, and function under future conditions.     

Copper and thermal perturbations on the early life processes of the hard coral <Emphasis Type="Italic">Platygyra acuta</Emphasis>

Anthropogenic pollutants and climate change are major threats to coral reefs today. Yet interactions between chemical and thermal perturbations have not been fully explored in reef studies. Here, we present the single and combined effects of copper (Cu) with thermal stress on five early life-history stages/processes (fertilization, larval mortality, swimming ability, metamorphosis and growth of juvenile recruits) of the massive coral Platygyra acuta in Hong Kong. In the first four experiments, coral gametes and larvae were exposed to different Cu doses (0–200 μg L^?1, apart from the fertilization assay in which 0–1000 μg L^?1 was used) and temperature treatments (ambient and ambient +2 or +3 °C as a thermal stress treatment) following a factorial experimental design. Exposure time was 5 h for the fertilization assay and 48 h for the other experiments. The last experiment on growth of coral recruits was conducted over 56 d with 0–80 μg L^?1 Cu used. Cu significantly reduced percent fertilization success, percentage of active swimming larvae and larval survivorship (EC₅₀s, the half maximal effective concentrations, for percent fertilization success and percentage of active swimming larvae were 92–145 and 45–47 μg L^?1 respectively. While LC₅₀, the lethal concentration that kills 50% of the population, was 101–110 μg L^?1), while growth of coral recruits was not affected at 80 μg L^?1 Cu for 56 d. No settling cues were used in the settlement experiment. In their absence, percent metamorphosis increased with Cu doses, in sharp contrast to earlier findings. Settlement and metamorphosis may thus be strategies for coral larvae to escape from Cu toxicity. Thermal treatment did not significantly affect any experimental end points. This is likely because the thermal regimes used in the experiments were within the range experienced by local corals. The high variability in Cu toxicities indicates differential susceptibilities of the various life-history stages/processes of P. acuta. The level of Cu tolerance was also markedly higher than that reported in the literature for other coral species. This provides evidence to suggest possible adaptation of this species to survive in a highly polluted marine environment like that in Hong Kong.     

Macroalgal herbivory on recovering versus degrading coral reefs

Macroalgal-feeding fishes are considered to be a key functional group on coral reefs due to their role in preventing phase shifts from coral to macroalgal dominance, and potentially reversing the shift should it occur. However, assessments of macroalgal herbivory using bioassay experiments are primarily from systems with relatively high coral cover. This raises the question of whether continued functionality can be ensured in degraded systems. It is clearly important to determine whether the species that remove macroalgae on coral-dominated reefs will still be present and performing significant algal removal on macroalgal-dominated reefs. We compared the identity and effectiveness of macroalgal-feeding fishes on reefs in two conditions post-disturbance—those regenerating with high live coral cover (20–46 %) and those degrading with high macroalgal cover (57–82 %). Using filmed Sargassum bioassays, we found significantly different Sargassum biomass loss between the two conditions; mean assay weight loss due to herbivory was 27.9 ± 4.9 % on coral-dominated reefs and 2.2 ± 1.1 % on reefs with high macroalgal cover. However, once standardised for the availability of macroalgae on the reefs, the rates of removal were similar between the two reef conditions (4.8 ± 4.1 g m^?2 h^?1 on coral-dominated and 5.3 ± 2.1 g m^?2 h^?1 on macroalgal-dominated reefs). Interestingly, the Sargassum-assay consumer assemblages differed between reef conditions; nominally grazing herbivores, Siganus puelloides and Chlorurus sordidus, and the browser, Siganus sutor, dominated feeding on high coral cover reefs, whereas browsing herbivores, Naso elegans, Naso unicornis, and Leptoscarus vaigiensis, prevailed on macroalgal-dominated reefs. It appeared that macroalgal density in the surrounding habitat had a strong influence on the species driving the process of macroalgal removal. This suggests that although the function of macroalgal removal may continue, the species responsible may change with context, differing between systems that are regenerating versus degrading.     

The effects of top–down versus bottom–up control on benthic coral reef community structure

While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral–algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.     

Sponge erosion under acidification and warming scenarios: differential impacts on living and dead coral

Ocean acidification will disproportionately impact the growth of calcifying organisms in coral reef ecosystems. Simultaneously, sponge bioerosion rates have been shown to increase as seawater pH decreases. We conducted a 20‐week experiment that included a 4‐week acclimation period with a high number of replicate tanks and a fully orthogonal design with two levels of temperature (ambient and +1 °C), three levels of pH (8.1, 7.8, and 7.6), and two levels of boring sponge (Cliona varians, present and absent) to account for differences in sponge attachment and carbonate change for both living and dead coral substrate (Porites furcata). Net coral calcification, net dissolution/bioerosion, coral and sponge survival, sponge attachment, and sponge symbiont health were evaluated. Additionally, we used the empirical data from the experiment to develop a stochastic simulation of carbonate change for small coral clusters (i.e., simulated reefs). Our findings suggest differential impacts of temperature, pH and sponge presence for living and dead corals. Net coral calcification (mg CaCO₃ cm^?2 day^?1) was significantly reduced in treatments with increased temperature (+1 °C) and when sponges were present; acidification had no significant effect on coral calcification. Net dissolution of dead coral was primarily driven by pH, regardless of sponge presence or seawater temperature. A reevaluation of the current paradigm of coral carbonate change under future acidification and warming scenarios should include ecologically relevant timescales, species interactions, and community organization to more accurately predict ecosystem‐level response to future conditions.     

Differential modification of seawater carbonate chemistry by major coral reef benthic communities

Ocean acidification (OA) resulting from uptake of anthropogenic CO₂ may negatively affect coral reefs by causing decreased rates of biogenic calcification and increased rates of CaCO₃ dissolution and bioerosion. However, in addition to the gradual decrease in seawater pH and Ω _a resulting from anthropogenic activities, seawater carbonate chemistry in these coastal ecosystems is also strongly influenced by the benthic metabolism which can either exacerbate or alleviate OA through net community calcification (NCC = calcification – CaCO₃ dissolution) and net community organic carbon production (NCP = primary production ? respiration). Therefore, to project OA on coral reefs, it is necessary to understand how different benthic communities modify the reef seawater carbonate chemistry. In this study, we used flow-through mesocosms to investigate the modification of seawater carbonate chemistry by benthic metabolism of five distinct reef communities [carbonate sand, crustose coralline algae (CCA), corals, fleshy algae, and a mixed community] under ambient and acidified conditions during summer and winter. The results showed that different communities had distinct influences on carbonate chemistry related to the relative importance of NCC and NCP. Sand, CCA, and corals exerted relatively small influences on seawater pH and Ω _a over diel cycles due to closely balanced NCC and NCP rates, whereas fleshy algae and mixed communities strongly elevated daytime pH and Ω _a due to high NCP rates. Interestingly, the influence on seawater pH at night was relatively small and quite similar across communities. NCC and NCP rates were not significantly affected by short-term acidification, but larger diel variability in pH was observed due to decreased seawater buffering capacity. Except for corals, increased net dissolution was observed at night for all communities under OA, partially buffering against nighttime acidification. Thus, algal-dominated areas of coral reefs and increased net CaCO₃ dissolution may partially counteract reductions in seawater pH associated with anthropogenic OA at the local scale.